large mammals from new late miocene fossil … medium sized hipparion species of maragheh are...
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LARGE MAMMALS FROM NEW LATE MIOCENE FOSSIL
LOCALITIES NEAR VARZEGHAN, NORTHWEST IRAN
Majid MIRZAIE ATAABADI1, Gholamreza ZAREE2, Zahra ORAK2
1 Department of Geosciences and Geography, University of Helsinki, P.O. Box 64,
FI-00014, Helsinki, Finland
2 Environment Protection Organization, MMTT, Tehran, Iran
Abstract
Large mammals from localities near Varzeghan in northwestern Iran are described here for the first
time. The fossil material consists of hipparionine horses and deinotheres. The hipparion species are
tentatively differentiated according to their size and dental morphology into a small to medium sized
species like Hipparion moldavicum and a medium to large sized species such as H. prostylum or H.
dietrichi. Deinotherium giganteum is also present in this area. A Middle Turolian age (MN 12
equivalent) is proposed for these new localities based on comparison of the fossil material to material
from other Late Miocene sites.
Key words Mammals, Miocene, Turolian, Varzeghan, Maragheh, Iran
1 Introduction Although extensive outcrops of Neogene
terrestrial deposits exist in northwest and
central Iran, knowledge of mammalian fossil
faunas in the country remains limited due to
lack of sufficient investigations. The only
exception is the richly fossiliferous beds of the
Maragheh Formation in northwest Iran. These
fossiliferous deposits are situated on the
southern and eastern slopes of the Sahand
volcano in the Maragheh district of eastern
Azarbaijan Province. For more than one and a
half centuries, this famous Late Miocene
vertebrate locality received much attention and
has been the exclusive source of information
on Neogene mammalian faunas of Iran. Based
on the latest studies the Maragheh Fm. has
been divided into three biostratigraphic
intervals (Lower, Middle and Upper
Maragheh), ranging from 9.5 to 7 million years
(Ma) in age (Bernor, 1986).
Nevertheless, recent discovery of new
localities in the northwestern parts of Iran has
demonstrated a wider distribution of mammal
bearing deposits in this area. These new
fossiliferous areas are situated north and north
east of Tabriz, the capital city of East
Azarbaijan province and are ca. 110-150 km
away from Maragheh (Fig. 1). The mammalian
fauna of Ivand, one of these new localities, has
been studied recently (Sen and Purabrishemi
2010, Mirzaie Ataabadi et al. ms). Here we
report on large mammal fossils from new
localities near Varzeghan.
MIRZAIE ATAABADI ET AL. / LATE MIOCENE LARGE MAMMALS FROM VARZEGHAN
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Fig. 1 The geographic location of the new late Miocene mammal fossil localities (Varzeghan and
Ivand) in NW Iran (map modified after Bernor, 1986)
2 Geological setting
During the Paleogene, Azarbaijan province in
northwest Iran experienced a wide range of
volcanic activities. By the end of the early
Miocene the last Tethyan seaway disappeared
from the area, carbonate deposition terminated,
and a major landmass emerged (Aghanabati,
2004). Consequently, at the beginning of the
Neogene, this domain was mostly above sea
level and had incipient mountain ranges, basin
troughs, and a topography resembling present
day conditions (Davoudzadeh et al., 1997).
The areas north of the Tabriz fault (the
Anatolian transform of figure 1) in particular,
are segments of the Tabriz-Maku sub-zone in
the western Alborz-Azarbaijan structural zone.
During the Late Miocene and Pliocene this
zone, like other parts of Iran, underwent major
orogenic movements that established new
erosional cycles that filled the locally closed
basins with fluvial and lacustrine deposits
(Aghanabati, 2004). The new fossil localities
in the Varzeghan (and Ivand) districts and their
mammalian faunas, bounded by the Anatolian
transform to the south (Fig. 1), were formed in
such local basin troughs with different
environments compared to Maragheh
Formation.
3 Fossil localities and material
Varzeghan is situated (ca. 60 km) northeast of
Tabriz (Fig. 1). This locality is less well known
than the Ivand locality, mainly because of its
more recent discovery as a consequence of
construction activities in the area. Because of
the occurrence of fossils on the surface and
lack of sediment exposures, the nature of the
fossil bearing sequence and especially its
sedimentology and depositional environment is
not well understood. In Abkhare village (ca
MIRZAIE ATAABADI ET AL. / LATE MIOCENE LARGE MAMMALS FROM VARZEGHAN
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38° 31' N, 46° 32' E), the fossil site of
deinothere material is a local concentration of
bones within a mudstone bed. The fossil
pocket is restricted to a single horizon.
Unfortunately, due to lack of outcrops
only a two-meter deep trench in Abkhare
village near the fossil site is the source of
sedimentology information. Here, fine grained
sediments are mainly present with a minor
lenticular pebbly sand layer (channel deposit)
which indicates a fluvial environment.
The deinothere material is partly
articulated. Fossils are very well preserved and
do not show any sign of weathering or
abrasion. The hipparion material is also very
well preserved. It seems that partly articulated
hipparions, as well as other fossil animals,
could be found at this locality in the future.
Therefore, the taphonomy of the fossil material
in this locality seems to be different from that
of Maragheh.
Fossil bones in Maragheh Fm. occur as
localized concentrations within the
unlaminated beds, floating in the sediments
rather than lying on bedding-planes. A single
complete articulated skeleton of the mustelid
Promeles palaeattica has been found from the
MMTT 13 quarry (Bernor et al., 1996).
Taphonomic studies of these fossil
accumulations indicate autochthonous bone
assemblages accumulated on overbank or
floodplain deposits of fluvial systems. A large
number of the bones are preserved with
articulation of distal limb elements and early
weathering stages. Pyroclastic events such as
mudflows or ash falls were not directly
responsible for mortality. On the other hand,
biologic agents were the probable cause of
death, as the bones were subaerially exposed
only long enough to allow removal of some
elements by scavengers (Morris, 1997).
The collected fossil material includes several
cranial and postcranial elements of a large
deinothere, which were unearthed while
renovating an old building in the outlying parts
of Abkhare village. These materials are stored
temporarily in the Department of Geology,
University of Tabriz, and the Department of
Environment local branch office in Maragheh.
In addition, about 100 isolated maxillary and
mandibular cheek teeth, 10 mandibular
fragments, part of a juvenile skull and a few
distal metapodials of hipparionine horses were
recovered from Varzeghan district. This
material is stored in the Department of
Environment local branch office in Maragheh.
The hipparion material has been
apparently collected from a construction site in
the city of Varzeghan (ca. 38o 30' N, 46
o 40'
E). However, we cannot exclude the possibility
of its mixture with fossils from other localities
in Varzeghan district.
The current study is based on the best
preserved mandibular fragments of hipparions
and the maxillary molar teeth of the
deinothere. Postcranial remains of the
deinothere were unavailable for this study. The
absence of other fossil groups such as bovids is
mostly due to collection bias and the limited
time available for collecting in the field.
4 Abbreviations
MMTT Muze Melli Tarikh Tabiei (National
Museum of Natural History), Tehran
MN European Mammal Neogene Faunal
Zone
M1-3 Upper Molars
p2-4 Lower premolars
m1-3 Lower molars
5 Systematic paleontology
Order Perissodactyla Owen, 1848
Family Equidae Gray, 1821
Genus Hipparion de Christol, 1832
Hipparion sp. 1 (small to medium sized)
Locality Varzeghan District
Age Middle Turolian, MN 12 equivalent
Material Right mandibular fragment with p2-
m3, MMTT-2-102-V318 (Fig 2-A, Tables 1-2)
Description The tooth row is relatively short
and the teeth are small. The premolars are wide
and robust. The length of p2-m3 is 130.8 mm
and the ratio (premolars length) ×100 / (molars
length) is 115, indicating a short molar row.
The parastylid is relatively well developed.
The metaconid and the metastylid are round to
subtriangular. The entoconid is round to
elliptical. The enamel is plicated at the flexid
borders, particularly in the premolars. The
ectoflexid is V-shaped and is moderately deep
and narrow on p4-m2, reaching the middle of
the tooth but not touching the linguaflexid. It is
deeper on m1-m2 but does not separate the
pre- and post-flexids. The linguaflexid is V-
shaped but shallower and more open on p3-p4.
The hypoconulid is well developed. A weak
plicabalinid is present on p2-p3.
Hipparion sp. 2 (medium to large sized)
Locality Varzeghan District
Age Middle Turolian, MN 12 equivalent
Material Left mandibular fragment with p3-
m3, MMTT-2-108-V324 (Fig. 2-F, Tables 1-
2); left mandibular fragment with p3-m3,
MMTT-2-105-V321 (Fig. 2-G, Tables 1-2);
right mandibular fragment with p3-m3 and the
symphysis, MMTT-2-101-V317 (Fig. 2C-D,
Tables 1-2); right mandibular fragment with
p2-m2, MMTT-2-106-V322 (Fig. 2-E, Tables
1-2); right mandibular fragment with p3-m1,
MMTT-2-103-V319 (Fig. 2-B, Tables 1-2);
right mandibular fragment with p3-m2,
MMTT-2-104-V320 (Fig. 2-H, Tables 1-2)
Description
The tooth row is relatively long and the teeth
are moderately large. The premolars are wide
and robust. The length of p2-m3 is estimated to
be 145-150 mm and the ratio (premolars
length) ×100 / (molars length) is estimated
114, indicating a short molar row. The
parastylid is well developed and closed. The
metaconid and the metastylid are round to
subtriangular. The entoconid is elliptical to
round. The enamel is plicated at the flexids and
their borders, particularly on the premolars.
The ectoflexid is U-shaped and moderately
deep and narrow on the premolars, reaching
the middle of the tooth but not touching the
linguaflexid and not separating pre- and post-
flexids. The ectoflexid is V-shaped and deep
on the molars, reaching the linguaflexid and
separating the pre- and post-flexids. The
linguaflexid is U-shaped, shallow and open. It
is deeper and V-shaped on the worn teeth. The
hypoconulid is well developed. A plicabalinid
is present especially on premolars.
TABLE 1 Measurements of Hipparion sp. 1 and sp. 2 mandibular fragments, Varzeghan District, NW
Iran
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Fig. 2 Hipparion sp. 1 and sp. 2, Varzeghan District, NW Iran. Mandibular fragment occlusal views:
A, V318: right p2-m3; B, V319: Right p3-m1; C-D, V317: right p3-m3; E, V322: right p2-m2; F, V324:
Left p3-m3; G, V321: left p3-m3; H, V320: Right p3-m2. Scale bars: 20 mm
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TABLE 2 Measurements of Hipparion sp. 1 and sp. 2 mandibular cheek teeth, Varzeghan District,
NW Iran
Comparisons
Two species of hipparionine horses from
Varzeghan are discriminated here principally
based on their size and dental morphology.
Hipparion sp. 1 is represented by a single
mandibular fragment and belongs to a small to
medium sized species while the rest of
mandibular fragments belong to a larger
species. The dental morphology of the
mandibular cheek teeth of these two species is
also somewhat different. The ectoflexid in the
larger species is deeper and the linguaflexid is
more open and shallow.
Due to the geographical proximity of the
Varzeghan and Maragheh localities,
comparison between hipparion materials of
these localities is of interest. The small and
medium sized hipparion species of Maragheh
are Hipparion matthewi and H. moldavicum.
The type material of the small hipparion H.
matthewi is from Samos Island in Greece. The
length of the lower tooth row of this species in
the type specimen is about 110mm. H.
moldavicum is the most abundant medium
sized hipparion species at Maragheh.
MIRZAIE ATAABADI ET AL. / LATE MIOCENE LARGE MAMMALS FROM VARZEGHAN
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It is also very widespread in the northern
Black sea region where the type specimen was
found. The upper tooth row of this species
from Maragheh has a length of 120-140 mm
(Bernor 1985). Also, a recent comprehensive
study on the equid material from the
Akkaşdaği locality in Turkey has documented
a range of 128-142 mm for the length of the
lower tooth row (Koufos and Vlachou, 2005)
of H. moldavicum. Therefore, according to
these data Hipparion sp. 1 from Varzeghan is
in the size range of H. moldavicum and might
belong to it. Features in the mandibular dental
morphology such as deeper ectoflexids and V-
shaped linguaflexids in the molars of
Hipparion sp. 1 from Varzeghan support this
assignment.
The medium to large sized hipparionine
horses from Maragheh are H. prostylum, H.
campbelli, and H. gettyi. H. prostylum is more
abundant and smaller in size than the other
species. The upper tooth row of this species is
generally 140-150 mm long (Bernor, 1985).
Koufos and Vlachou (2005) have pointed to
the close similarity of H. dietrichi from
Akkaşdaği and some Greek localities to H.
prostylum from Maragheh. They recorded a
length of 138-145 mm for the lower cheek
tooth row of H. dietrichi from the Akkaşdaği
locality in Turkey. Thus, Hipparion sp. 2 from
Varzeghan is similar in size to H. prostylum
and H. dietrichi and might belong to either of
these species. Similarity in mandibular cheek
tooth morphology between these species, such
as deep ectoflexids and shallow and open
linguaflexids and a weak plicabalinid support
this idea. Unfortunately the muzzle is not
preserved in most of these fossils and hence
the main characteristic of H. dietrichi-H.
prostylum, which is the short and wide snout,
cannot be determined. Only one specimen
(MMTT-2-101-V317, Fig. 2-C) has the
symphysis partly preserved. In this individual
the short diastema is similar to that of H.
dietrichi, suggesting a short snout.
Order Proboscidea Illiger, 1811
Suborder Deinotherioidea Osborn, 1921
Family Deinotheriidae Bonaparte, 1845
Genus Deinotherium Kaup, 1829
Deinotherium giganteum Kaup, 1829
Locality Abkhare village, Varzeghan District
Age Middle Turolian, MN 12 equivalent
Material Upper molars including right M1,
MMTT-V16; M2, MMTT-V11 and M3,
MMTT-V14 and left M1, MMTT-V12; M2,
MMTT-V15 and M3, MMTT-V13 (Fig. 3,
Table 3).
Description
M1 (Fig. 3A-B, Table 3): The teeth are
trilophodont and have three roots. The three
transverse lophids are almost the same size. A
prominent cingulum is present on the anterior
wall. The lophids are heavily worn and partly
damaged.
M2 (Fig. 3C-D, Table 3): The teeth are
bilophodont and nearly quadratic in occlusal
outline and have three roots. The two
transverse lophids are of equal width. The teeth
are worn and damaged; nevertheless the labial
and posterior cingulae are clear. Both the
protoloph and metaloph have protocristids.
M3 (Fig. 3E-F, Table 3): The teeth are
bilophodont and have three roots. Right M3 is
very well preserved and has cingulum on the
anterior, labial, and posterior walls. The
protoloph is narrower than the metaloph and
has a less pronounced protocrista with a medial
extension.
Fig. 3 Deinotherium giganteum, Abkhare village, Varzeghan District, NW Iran. Upper molars,
occlusal views. A, V16: right M1 and B, V12: left M1. Scale bar: 40mm; C, V11: right M2 and D, V15:
left M2. Scale bar: 21 mm; E, V14: right M3 and F, V13: left M3. Scale bar: 21mm
TABLE 3 Measurements of Deinotherium giganteum upper Molars, Abkhare village, Varzeghan
District, NW Iran
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Comparison
The basic morphologies of the molar teeth of
species of Deinotheriidae are similar and the
main differences are in their size. Previous
studies have shown that the size of deinotheriid
dental and skeletal elements increased
gradually throughout the Miocene (Huttunen,
2002). The Deinotherium material from
Abkhare belongs to a single and very large
individual. Figure 4 shows the bivariate plots
of length and width of the molar teeth of the
Varzeghan specimens and other material from
Middle to Late Miocene localities in Europe.
The size of the teeth of Deinotherium from
Varzeghan as figured in the plots is larger than
the material from Pontlevoy (MN5), Sansan
(MN 6) and Montredon (MN10) in France,
Deinotheriensande and Eppelsheim (MN 9) in
Germany, Pannonian basin in central Europe
(MN7-10) and Late Miocene localities in
Greece, Bulgaria and Hungary. Although
slightly smaller, the Deinotherium molars from
Varzeghan are comparable in size to the teeth
of Deinotherium giganteum from Obuhovka
(MN 12) of southern Russia (Bajgusheva and
Titov, 2006), but are smaller than teeth of D.
gigantissimum.
6 Biochronology and paleoenvironment
Although the new fossil localities at
Varzeghan (and Ivand) are separated from the
Maragheh Fm. by the Anatolian transform
(Fig.1), but because of their geographical
proximity (ca. 110-150 kilometres), faunal
comparison and correlation of these fossil sites
are of interest. Like other typical Turolian
localities of the old world, hipparions are the
most abundant material in the Varzeghan
district.
Among the known hipparionine horses
from Maragheh medium sized hipparion
species such as Hipparion moldavicum and
medium to large sized species such as H.
prostylum are the most common species. Both
these species occur in the Middle Maragheh
biostratigraphical interval, which is most likely
8 to 7 million years old (Bernor, 1985; 1986).
Species of similar size and related to H.
prostylum, such as H. dietrichi, have a wide
distribution in the eastern Mediterranean
localities and are recorded from many
localities in the Balkans and Anatolia.
Occurrence of this latter species is mostly in
MN12 (8-7 Ma), but it is also reported from
MN 11 localities. H. moldavicum is also very
common in the northern Black sea region and
is recorded from MN11-12 while being absent
from the Balkans (Koufos and Vlachou, 2005).
The only occurrence of H. moldavicum in
Anatolia is from Akkaşdaği at 7.1 Ma (Koufos
and Vlachou, 2005).
The Ivand locality is another new Late
Miocene fossil site in northwest Iran. The
Varzeghan district is closer to this locality (ca.
50 kilometers) than to Maragheh. Hipparionine
horses are also abundant in this area, but only a
large sized species very similar to H.
brachypus is recorded from Ivand and the rest
of the material has not been studied yet. The
occurrence of H. brachypus at Pikermi and
Samos (Greece), Akkaşdaği (Turkey), and
Hadjidimovo (Bulgaria) is in the middle
Turolian or MN12 (Koufos and Vlachou,
2005). Consequently a Middle Turolian age
(ca. 8-7 Ma) can be inferred for the levels
containing Hipparion sp. large at Ivand. This is
also consistent with the age span of the Group
1 hipparionine horses of Bernor (1985).
Although Deinotherium giganteum has a
broad stratigraphical range from the Middle
Miocene to the Pliocene (18-2 Ma), given the
deinotheriid trend toward increasing size
during the Miocene (Huttunen, 2002) and the
very large size of the deinotheriid material
from Varzeghan, a late Miocene age of this
locality is evident.
MIRZAIE ATAABADI ET AL. / LATE MIOCENE LARGE MAMMALS FROM VARZEGHAN
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Based on the occurrence of
Deinotherium at Varzeghan, the locality can be
correlated with the Upper Maragheh
biostratigraphical interval, since the only
known Deinotherium from the Maragheh
Formation is recorded from the upper intervals
of this formation (K1 locality of Erdbrink et
al., 1976; identical to MMTT locality 31 of
Bernor, 1986). Absolute ages obtained from
Zircon fission track and K/Ar dating of
pumicites from some higher levels of the
Maragheh Formation implies an age of 7.4 Ma
for the uppermost part of this formation
(Bernor, 1986), which can be also assigned to
the Deinotherium-yielding upper levels of the
Maragheh Formation. D. giganteum is also
represented by several postcranial elements
from the Ivand locality. Based on the presence
of H. brachypus at this locality a Middle
Turolian age has been proposed for the
locality.
In conclusion, comparisons of the fossil
material from Varzeghan with other localities
in northwest Iran and the chronological
occurrences of similar taxa, suggest a middle
Turolian age (MN 12 equivalent) for the fossil
sites near Varzeghan.
The morphology of the deinotheriid
cheek teeth, which are lophodont and
brachyodont, indicates a folivorous diet and
implies a forested environment. However, the
very large size of the animal suggests a
landscape more open than that of a forest
(Poulakakis et al. 2005). Therefore, the
presence of this species in the new Late
Miocene fossil localities in northwest Iran
indicates either woodland or an open forest as
the predominant environment.
7 Conclusions
The area near Varzeghan in northwest Iran has
produced several new fossil localities. As a
consequence, our knowledge of Late Miocene
mammals in Iran is no longer restricted to the
fauna of the Maragheh Formation.
Hipparionine horses and deinotheres are
among the fossil material so far collected from
this area. The study of mandibular fragments
of hipparionine horses has resulted in
identification of two hipparion species based
on size and dental morphology. These fossil
materials belong to small to medium sized
hipparion species such as Hipparion
moldavicum and to medium to large sized
species such as H. prostylum or H. dietrichi.
According to the age and distribution of
these latter taxa in the eastern Mediterranean
and northern Black Sea regions and correlation
of the new localities with Maragheh and Ivand,
a Middle Turolian age (MN 12 equivalent) is
proposed for the sites near Varzeghan. The
presence of Deinotherium giganteum is very
important for correlation of these new
localities to biostratigraphical intervals of the
Maragheh Formation and Ivand and has proved
the existence of semi-open environments in the
region. More systematic collection of fossils at
Varzeghan and study of additional material
such as the maxillary cheek teeth of the
hipparionine horses will enhance our
understanding of these new localities and their
relationships with contemporaneous faunas in
northwest Iran and the adjacent Caucasus
region.
Acknowledgements
Iran’s Environment Protection Organization
(formerly Department of Environment)
facilitated this research and the field work. We
appreciate the pioneering work of Dr Z.
Purabrishemi, Tabriz University, in the
Varzeghan district. We are also grateful to
Prof. J. Barry, Peabody Museum, for
improving the text of this paper and his
constructive comments. MMA thanks the
support from Academy of Finland and RHOI
project.
MIRZAIE ATAABADI ET AL. / LATE MIOCENE LARGE MAMMALS FROM VARZEGHAN
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Fig. 4 Scatter plots of M1, M2 and M3 length against width in Deinotherium giganteum from Abkhare
village, Varzeghan District, and deinotheres from twelve European localities (data from Huttunen,
2002; Poulakakis et al., 2005; Bajgusheva and Titov, 2006)
× Deinotherium giganteum, Varzeghan; - D. giganteum, Sansan (MN6); + D. giganteum, Eppelsheim
(MN9); D. giganteum, Hungary (L. Miocene); D. giganteum, Obuhovka (MN11); ▲ D.
giganteum, Pontlevoy (MN5); Δ Prodeinotherium, Austria (Pannonian); D. giganteum, Siteia
(MN11); D. giganteum, Bulgaria (L. Miocene); ● D. giganteum, Deinotheriensande (MN9); ○ D.
giganteum, Montredon (MN10); ■ D. giganteum Austria (MN7-10); □ D. gigantissimum, Romania (L.
Miocene)
MIRZAIE ATAABADI ET AL. / LATE MIOCENE LARGE MAMMALS FROM VARZEGHAN
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